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Cnidaria () is a phylum under kingdom Animalia containing over 11,000
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate ...
of
aquatic animal An aquatic animal is any animal, whether invertebrate or vertebrate, that lives in water for most or all of its lifetime. Many insects such as mosquitoes, mayflies, dragonflies and caddisflies have aquatic larvae, with winged adults. Aquatic anim ...
s found both in freshwater and marine environments, predominantly the latter. Their distinguishing feature is
cnidocyte A cnidocyte (also known as a cnidoblast or nematocyte) is an explosive cell containing one large secretory organelle called a cnidocyst (also known as a cnida () or nematocyst) that can deliver a sting to other organisms. The presence of this c ...
s, specialized cells that they use mainly for capturing prey. Their bodies consist of mesoglea, a non-living jelly-like substance, sandwiched between two layers of epithelium that are mostly one cell thick. Cnidarians mostly have two basic body forms: swimming
medusa In Greek mythology, Medusa (; Ancient Greek: Μέδουσα "guardian, protectress"), also called Gorgo, was one of the three monstrous Gorgons, generally described as winged human females with living venomous snakes in place of hair. Those ...
e and sessile polyps, both of which are radially symmetrical with mouths surrounded by
tentacle In zoology, a tentacle is a flexible, mobile, and elongated organ present in some species of animals, most of them invertebrates. In animal anatomy, tentacles usually occur in one or more pairs. Anatomically, the tentacles of animals work mainl ...
s that bear cnidocytes. Both forms have a single orifice and body cavity that are used for
digestion Digestion is the breakdown of large insoluble food molecules into small water-soluble food molecules so that they can be absorbed into the watery blood plasma. In certain organisms, these smaller substances are absorbed through the small intes ...
and respiration. Many cnidarian species produce
colonies In modern parlance, a colony is a territory subject to a form of foreign rule. Though dominated by the foreign colonizers, colonies remain separate from the administration of the original country of the colonizers, the '' metropolitan state'' ...
that are single organisms composed of medusa-like or polyp-like zooids, or both (hence they are trimorphic). Cnidarians' activities are coordinated by a decentralized nerve net and simple receptors. Several free-swimming species of
Cubozoa Box jellyfish (class Cubozoa) are cnidarian invertebrates distinguished by their box-like (i.e. cube-shaped) body. Some species of box jellyfish produce potent venom delivered by contact with their tentacles. Stings from some species, including '' ...
and Scyphozoa possess balance-sensing
statocyst The statocyst is a balance sensory receptor present in some aquatic invertebrates, including bivalves, cnidarians, ctenophorans, echinoderms, cephalopods, and crustaceans. A similar structure is also found in ''Xenoturbella''. The statocyst con ...
s, and some have simple eyes. Not all cnidarians reproduce sexually, but many species have complex life cycles of asexual polyp stages and sexual medusae stages. Some, however, omit either the polyp or the medusa stage, and the parasitic classes evolved to have neither form. Cnidarians were formerly grouped with
ctenophore Ctenophora (; ctenophore ; ) comprise a phylum of marine invertebrates, commonly known as comb jellies, that inhabit sea waters worldwide. They are notable for the groups of cilia they use for swimming (commonly referred to as "combs"), and ...
s in the phylum
Coelenterata Coelenterata is a term encompassing the animal phyla Cnidaria (coral animals, true jellies, sea anemones, sea pens, and their relatives) and Ctenophora (comb jellies). The name comes , referring to the hollow body cavity common to these two ph ...
, but increasing awareness of their differences caused them to be placed in separate phyla. Cnidarians are classified into four main groups: the almost wholly sessile
Anthozoa Anthozoa is a subphylum of marine invertebrates which includes the sea anemones, stony corals and soft corals. Adult anthozoans are almost all attached to the seabed, while their larvae can disperse as part of the plankton. The basic unit o ...
(
sea anemone Sea anemones are a group of predatory marine invertebrates of the order Actiniaria. Because of their colourful appearance, they are named after the '' Anemone'', a terrestrial flowering plant. Sea anemones are classified in the phylum Cnidaria ...
s, corals, sea pens); swimming Scyphozoa ( jellyfish);
Cubozoa Box jellyfish (class Cubozoa) are cnidarian invertebrates distinguished by their box-like (i.e. cube-shaped) body. Some species of box jellyfish produce potent venom delivered by contact with their tentacles. Stings from some species, including '' ...
(box jellies); and Hydrozoa (a diverse group that includes all the freshwater cnidarians as well as many marine forms, and has both sessile members, such as '' Hydra'', and colonial swimmers, such as the Portuguese Man o' War). Staurozoa have recently been recognised as a
class Class or The Class may refer to: Common uses not otherwise categorized * Class (biology), a taxonomic rank * Class (knowledge representation), a collection of individuals or objects * Class (philosophy), an analytical concept used differentl ...
in their own right rather than a sub-group of Scyphozoa, and the highly derived parasitic
Myxozoa Myxozoa (etymology: Greek: μύξα ''myxa'' "slime" or "mucus" + thematic vowel o + ζῷον ''zoon'' "animal") is a subphylum of aquatic cnidarian animals – all obligate parasites. It contains the smallest animals ever known to have lived. ...
and Polypodiozoa were firmly recognized as cnidarians in 2007. Most cnidarians prey on organisms ranging in size from plankton to animals several times larger than themselves, but many obtain much of their nutrition from dinoflagellates, and a few are parasites. Many are preyed on by other animals including
starfish Starfish or sea stars are star-shaped echinoderms belonging to the class Asteroidea (). Common usage frequently finds these names being also applied to ophiuroids, which are correctly referred to as brittle stars or basket stars. Starfish ...
, sea slugs,
fish Fish are Aquatic animal, aquatic, craniate, gill-bearing animals that lack Limb (anatomy), limbs with Digit (anatomy), digits. Included in this definition are the living hagfish, lampreys, and Chondrichthyes, cartilaginous and bony fish as we ...
,
turtle Turtles are an order of reptiles known as Testudines, characterized by a special shell developed mainly from their ribs. Modern turtles are divided into two major groups, the Pleurodira (side necked turtles) and Cryptodira (hidden necked ...
s, and even other cnidarians. Many
scleractinia Scleractinia, also called stony corals or hard corals, are marine animals in the phylum Cnidaria that build themselves a hard skeleton. The individual animals are known as polyps and have a cylindrical body crowned by an oral disc in which a mo ...
n corals—which form the structural foundation for coral reefs—possess polyps that are filled with symbiotic photo-synthetic
zooxanthellae Zooxanthellae is a colloquial term for single-celled dinoflagellates that are able to live in symbiosis with diverse marine invertebrates including demosponges, corals, jellyfish, and nudibranchs. Most known zooxanthellae are in the genus '' Symb ...
. While reef-forming corals are almost entirely restricted to warm and shallow marine waters, other cnidarians can be found at great depths, in
polar region The polar regions, also called the frigid zones or polar zones, of Earth are the regions of the planet that surround its geographical poles (the North and South Poles), lying within the polar circles. These high latitudes are dominated by fl ...
s, and in freshwater. Recent
phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
analyses support monophyly of cnidarians, as well as the position of cnidarians as the sister group of
bilateria The Bilateria or bilaterians are animals with bilateral symmetry as an embryo, i.e. having a left and a right side that are mirror images of each other. This also means they have a head and a tail (anterior-posterior axis) as well as a belly and ...
ns. Fossil cnidarians have been found in rocks formed about , and other fossils show that corals may have been present shortly before and diversified a few million years later. However,
molecular clock The molecular clock is a figurative term for a technique that uses the mutation rate of biomolecules to deduce the time in prehistory when two or more life forms diverged. The biomolecular data used for such calculations are usually nucleoti ...
analysis of
mitochondrial A mitochondrion (; ) is an organelle found in the cells of most Eukaryotes, such as animals, plants and fungi. Mitochondria have a double membrane structure and use aerobic respiration to generate adenosine triphosphate (ATP), which is used t ...
genes suggests a much older age for the
crown group In phylogenetics, the crown group or crown assemblage is a collection of species composed of the living representatives of the collection, the most recent common ancestor of the collection, and all descendants of the most recent common ancestor. ...
of cnidarians, estimated around , almost 200 million years before the
Cambrian period The Cambrian Period ( ; sometimes symbolized Ꞓ) was the first geological period of the Paleozoic Era, and of the Phanerozoic Eon. The Cambrian lasted 53.4 million years from the end of the preceding Ediacaran Period 538.8 million years ago ...
as well as any fossils.


Distinguishing features

Cnidarians form a phylum of animals that are more complex than
sponge Sponges, the members of the phylum Porifera (; meaning 'pore bearer'), are a basal animal clade as a sister of the diploblasts. They are multicellular organisms that have bodies full of pores and channels allowing water to circulate through ...
s, about as complex as
ctenophore Ctenophora (; ctenophore ; ) comprise a phylum of marine invertebrates, commonly known as comb jellies, that inhabit sea waters worldwide. They are notable for the groups of cilia they use for swimming (commonly referred to as "combs"), and ...
s (comb jellies), and less complex than
bilateria The Bilateria or bilaterians are animals with bilateral symmetry as an embryo, i.e. having a left and a right side that are mirror images of each other. This also means they have a head and a tail (anterior-posterior axis) as well as a belly and ...
ns, which include almost all other animals. Both cnidarians and ctenophores are more complex than sponges as they have: cells bound by inter-cell connections and carpet-like
basement membrane The basement membrane is a thin, pliable sheet-like type of extracellular matrix that provides cell and tissue support and acts as a platform for complex signalling. The basement membrane sits between epithelial tissues including mesothelium and ...
s;
muscle Skeletal muscles (commonly referred to as muscles) are organs of the vertebrate muscular system and typically are attached by tendons to bones of a skeleton. The muscle cells of skeletal muscles are much longer than in the other types of mus ...
s;
nervous system In biology, the nervous system is the highly complex part of an animal that coordinates its actions and sensory information by transmitting signals to and from different parts of its body. The nervous system detects environmental changes ...
s; and some have sensory organs. Cnidarians are distinguished from all other animals by having
cnidocyte A cnidocyte (also known as a cnidoblast or nematocyte) is an explosive cell containing one large secretory organelle called a cnidocyst (also known as a cnida () or nematocyst) that can deliver a sting to other organisms. The presence of this c ...
s that fire harpoon like structures and are usually used mainly to capture prey. In some species, cnidocytes can also be used as anchors. Cnidarians are also distinguished by the fact that they have only one opening in their body for ingestion and excretion i.e. they don't have a separate mouth and anus. Like sponges and ctenophores, cnidarians have two main layers of cells that sandwich a middle layer of jelly-like material, which is called the mesoglea in cnidarians; more complex animals have three main cell layers and no intermediate jelly-like layer. Hence, cnidarians and ctenophores have traditionally been labelled
diploblastic Diploblasty is a condition of the blastula in which there are two primary germ layers: the ectoderm and endoderm. Diploblastic organisms are organisms which develop from such a blastula, and include cnidaria and ctenophora, formerly grouped togeth ...
, along with sponges. However, both cnidarians and ctenophores have a type of
muscle Skeletal muscles (commonly referred to as muscles) are organs of the vertebrate muscular system and typically are attached by tendons to bones of a skeleton. The muscle cells of skeletal muscles are much longer than in the other types of mus ...
that, in more complex animals, arises from the middle cell layer. As a result, some recent text books classify ctenophores as
triploblastic Triploblasty is a condition of the gastrula in which there are three primary germ layers: the ectoderm, mesoderm, and endoderm. Germ cells are set aside in the embryo at the blastula stage, which are incorporated into the gonads during organoge ...
, and it has been suggested that cnidarians evolved from triploblastic ancestors.


Description


Basic body forms

Most adult cnidarians appear as either free-swimming
medusa In Greek mythology, Medusa (; Ancient Greek: Μέδουσα "guardian, protectress"), also called Gorgo, was one of the three monstrous Gorgons, generally described as winged human females with living venomous snakes in place of hair. Those ...
e or sessile polyps, and many hydrozoans species are known to alternate between the two forms. Both are radially symmetrical, like a wheel and a tube respectively. Since these animals have no heads, their ends are described as "oral" (nearest the mouth) and "aboral" (furthest from the mouth). Most have fringes of tentacles equipped with
cnidocyte A cnidocyte (also known as a cnidoblast or nematocyte) is an explosive cell containing one large secretory organelle called a cnidocyst (also known as a cnida () or nematocyst) that can deliver a sting to other organisms. The presence of this c ...
s around their edges, and medusae generally have an inner ring of tentacles around the mouth. Some hydroids may consist of colonies of zooids that serve different purposes, such as defense, reproduction and catching prey. The mesoglea of polyps is usually thin and often soft, but that of medusae is usually thick and springy, so that it returns to its original shape after muscles around the edge have contracted to squeeze water out, enabling medusae to swim by a sort of
jet propulsion Jet propulsion is the propulsion of an object in one direction, produced by ejecting a jet of fluid in the opposite direction. By Newton's third law, the moving body is propelled in the opposite direction to the jet. Reaction engines operating o ...
.


Skeletons

In medusae the only supporting structure is the mesoglea. '' Hydra'' and most
sea anemone Sea anemones are a group of predatory marine invertebrates of the order Actiniaria. Because of their colourful appearance, they are named after the '' Anemone'', a terrestrial flowering plant. Sea anemones are classified in the phylum Cnidaria ...
s close their mouths when they are not feeding, and the water in the digestive cavity then acts as a hydrostatic skeleton, rather like a water-filled balloon. Other polyps such as '' Tubularia'' use columns of water-filled cells for support. Sea pens stiffen the mesoglea with calcium carbonate spicules and tough fibrous
protein Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residues. Proteins perform a vast array of functions within organisms, including catalysing metabolic reactions, DNA replication, resp ...
s, rather like
sponge Sponges, the members of the phylum Porifera (; meaning 'pore bearer'), are a basal animal clade as a sister of the diploblasts. They are multicellular organisms that have bodies full of pores and channels allowing water to circulate through ...
s. In some colonial polyps, a chitinous periderm gives support and some protection to the connecting sections and to the lower parts of individual polyps. Stony corals secrete massive calcium carbonate exoskeletons. A few polyps collect materials such as sand grains and shell fragments, which they attach to their outsides. Some colonial sea anemones stiffen the mesoglea with
sediment Sediment is a naturally occurring material that is broken down by processes of weathering and erosion, and is subsequently transported by the action of wind, water, or ice or by the force of gravity acting on the particles. For example, sand ...
particles.


Main cell layers

Cnidaria are
diploblastic Diploblasty is a condition of the blastula in which there are two primary germ layers: the ectoderm and endoderm. Diploblastic organisms are organisms which develop from such a blastula, and include cnidaria and ctenophora, formerly grouped togeth ...
animals; in other words, they have two main cell layers, while more complex animals are triploblasts having three main layers. The two main cell layers of cnidarians form
epithelia Epithelium or epithelial tissue is one of the four basic types of animal tissue, along with connective tissue, muscle tissue and nervous tissue. It is a thin, continuous, protective layer of compactly packed cells with a little intercellula ...
that are mostly one cell thick, and are attached to a fibrous
basement membrane The basement membrane is a thin, pliable sheet-like type of extracellular matrix that provides cell and tissue support and acts as a platform for complex signalling. The basement membrane sits between epithelial tissues including mesothelium and ...
, which they secrete. They also secrete the jelly-like mesoglea that separates the layers. The layer that faces outwards, known as the ectoderm ("outside skin"), generally contains the following types of cells: *Epitheliomuscular cells whose bodies form part of the epithelium but whose bases extend to form
muscle Skeletal muscles (commonly referred to as muscles) are organs of the vertebrate muscular system and typically are attached by tendons to bones of a skeleton. The muscle cells of skeletal muscles are much longer than in the other types of mus ...
fibers in parallel rows. The fibers of the outward-facing cell layer generally run at right angles to the fibers of the inward-facing one. In
Anthozoa Anthozoa is a subphylum of marine invertebrates which includes the sea anemones, stony corals and soft corals. Adult anthozoans are almost all attached to the seabed, while their larvae can disperse as part of the plankton. The basic unit o ...
(anemones, corals, etc.) and Scyphozoa (jellyfish), the mesoglea also contains some muscle cells. *
Cnidocyte A cnidocyte (also known as a cnidoblast or nematocyte) is an explosive cell containing one large secretory organelle called a cnidocyst (also known as a cnida () or nematocyst) that can deliver a sting to other organisms. The presence of this c ...
s, the harpoon-like "nettle cells" that give the phylum Cnidaria its name. These appear between or sometimes on top of the muscle cells. * Nerve cells. Sensory cells appear between or sometimes on top of the muscle cells, and communicate via synapses (gaps across which chemical signals flow) with
motor nerve A motor nerve is a nerve that transmits motor signals from the central nervous system (CNS) to the muscles of the body. This is different from the motor neuron, which includes a cell body and branching of dendrites, while the nerve is made up of ...
cells, which lie mostly between the bases of the muscle cells. Some form a simple nerve net. *Interstitial cells, which are unspecialized and can replace lost or damaged cells by transforming into the appropriate types. These are found between the bases of muscle cells. In addition to epitheliomuscular, nerve and interstitial cells, the inward-facing gastroderm ("stomach skin") contains
gland In animals, a gland is a group of cells in an animal's body that synthesizes substances (such as hormones) for release into the bloodstream ( endocrine gland) or into cavities inside the body or its outer surface ( exocrine gland). Structure ...
cells that secrete digestive
enzyme Enzymes () are proteins that act as biological catalysts by accelerating chemical reactions. The molecules upon which enzymes may act are called substrates, and the enzyme converts the substrates into different molecules known as product ...
s. In some species it also contains low concentrations of cnidocytes, which are used to subdue prey that is still struggling. The mesoglea contains small numbers of
amoeba An amoeba (; less commonly spelled ameba or amœba; plural ''am(o)ebas'' or ''am(o)ebae'' ), often called an amoeboid, is a type of cell or unicellular organism with the ability to alter its shape, primarily by extending and retracting pseudo ...
-like cells, and muscle cells in some species. However, the number of middle-layer cells and types are much lower than in sponges.


Polymorphism

Polymorphism refers to the occurrence of structurally and functionally more than two different types of individuals within the same organism. It is a characteristic feature of Cnidarians, particularly the polyp and
medusa In Greek mythology, Medusa (; Ancient Greek: Μέδουσα "guardian, protectress"), also called Gorgo, was one of the three monstrous Gorgons, generally described as winged human females with living venomous snakes in place of hair. Those ...
forms, or of zooids within colonial organisms like those in Hydrozoa. In Hydrozoans, colonial individuals arising from individuals zooids will take on separate tasks. For example, in '' Obelia'' there are feeding individuals, the